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A Revision of Austmlopithecine Body Sizes

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A Revision of Austmlopithecine Body Sizes 65 A REVISION OF AUSTMLOPITHECINE BODY SIZES Grover S. Krantz Anthropology Department Washington State University Pullman, Washington 99L63, U.S.A. Received February 18, L977 ABSTMCT: When AustralopiEhecus dentiEions are divided into africanus and robustus on morphological grounds alone it is found that both species are found in each of the major South African sites. There is a greaL sexual dimor- phisrn in both species with a relative scarcity of the big urale africanus and the small female robustus. Skulls and jaws of young male africanus and female robustus have previously been misidentified as early genus Homg. With site allocation no longer valid, a ner^rmethod of identifying postcranial remains shows that africanus averages much larger than robustus. With new ratios of tooth fo body size the greater degree of morphological molarization in robustus now makes sense. J Introduction Australopithecus remains are generally thought of as two species; A. elg- ttgracilet' t'robusttt canus is the form, and A. robustus is the form. Some think this distinction merely reflects differences beLween sexes, races, and indivi- dual variations within a single species. Ot.hers raise the distinction to generic level with Paranghlgpge being the "robust't form, and Australopithecus continu- ing as tfre \ra"ifettorm which is sometimes even ittcl.rded in our own genus as Homo africanus. These various opinions are well known in the literature. The taxonomic leve1 of distinction between the trvo forms i-s not at issue here, but rather the most basic assumptions on which the two forms are based. Before anything meaningful can be decided about two kj-nds of australopithecines it is necessary to know which specimens belong in each category and how they differ from each other. The following discussj-on will center mainly on the South African specimens and extend from Lhere as pertinent. To most readers this is already well knownl robustus is the large form found at Swartkrans and Kromdraai, while africanus is the small form found at Taung, Sterkfontein, and Makapansgat. Other differences often discussed include molarization, cranial architecture, crest development, and locomotor adaptations. There is much disagreement about the nature, significance, or even the renli gy of these other differences, though everyone seems to agree they can be distinguished by sLze and by site. But apparently we have all been misled on both pointsl Two types of australopithecines can be distinguished in terms of dentitions, but it will be shown here that both of these occur mixed together in the vari-- ous sites. Since this would invalidate postcranial assignments based on the sites of recovery, other methods must be used for such assignments. The other methods developed here show that africanus is actually larger than robustus. This conclusion, if demonstrated, serves to clarify some problems, and may somewhat affect the phylogenies that are drawn up relating these forms to each other and to other hominids. It ls somewhat of a mystery to me how the australopithecine remains came Lo be classified according to site. Nowhere in the literature can one find a a:kt( Evol. Theory 2265-94 (May, L977) The editors thank two anonymous referees for help in evaluating this paper. @L977, The UniversitY of Chicago. 66 G. S. KRANTZ clear statement of how it is known that all specimens from a given South African cave deposit are of the same type. Somehave wondered about such site exclu- siveness, but that is about as far as it has gone. This arbitrary categorization has been uni-versally accepted by workers who have made comparisons between what they consider to be samples of the two forms. In all normal paleontology any pooling of specimens into a sample rests on the identification of every single individual as belonging to that taxon. Even in the australopithecine sites, other ani-mals are handled properly. If a few hyenas in one site are identified as belonging to a given species it is noE claimed that all hyenid remains from that site autornatically belong to that same species. It has not been expl-ained why the hominid remains were pooled without regard to demonstrating the affinities of each individual specimen. Some question has recently been raised that Makapansgat may contain speci- mens of both forms, or that it may represent a sample drawn from the point of time where they were only beginning to differentiate from each other (Tobias, L973). This idea is supported by observations on the fossils themselves (Wal- lace, L973, and Aguirre, 1970) and also by the strong possibility that Maka- pansgat may in fact be the earliest of all theaustralopithecinesites (We11sr1969). The universal- acceptance that robustus is larger in body size than africanus follows directly from this al-locatioi=byGite. Some hominid postcraniaT=66G- of fairly large size were found at Swartkrans and some very much smaller bones come from Sterkfontein. Since these constitute most of the hominid bones, and they correspond to the size contrast of rnost of the skull-s from these two sites, the body size contrast is easil-y postul-ated. The bones that donrt fit this picture, smaLl ones from Swartkrans and large ones from Sterkfontein, are either ignored or referred to other taxa entirely. In no case can cranial and post- cranial remains be related to the same individual-; mere physicaL proximity is not sufficient in these random bone accumul-ations. From here it is a simple matter to cl-assify any smal-l australopithecine bone as africanus, and every large one as robustus, wherever they are found. But if site alLoeation is not aLways correct, then the size distinction night not always be correct, and the ramificatlons get interesting. A11 of the evidence and reasoning behind these assignments needs to be re-examined with some care, and a modest beginning on this will be made here. Two Dental TYPes The usual pracLice has been to equate degree of molarizatLon with size and site. The more molarized dentitions go with larger body sizes and come from "robustn sites. In order to test the reality of this assumed correlation it is first necessary to divide the known dentitions into two morphologically distinct types without regard to their place of origin. This can be done rather easily for many speci-mens with the published measurements and descriptions. The length and breadth of each measurable tooth are here urultiplied together to give approximatlons of the cror^rn surface area. These are exaggerated because the teeth are not rectangular, but the exagerat.ionis consistent in all specimens. These surface areas are then arranged in the form of a bar graph for each jaw, beginning with the first incisor on the left and running to the third molar. The line connecting the tops of these area measurements constitutes what rnight be called a dental profile. So far this procedure is not greatly different from that used by Robinson in manypublications to illustrate the sequence of sizes of australopitheeine teeth. One minor difference is that instead of area he used dental module, the mean of length and breadth, which relatively exaggerates those teeth that are more nearly square. 67 AUSTMLOPITHECINEBODY SIZES A major difference is that Robinson, like all others, made his definitive comparisons between the dental profiles of pooled samples. Means and ranges of "robust" specimens were conLrasted with those of 'tgracileil specimens on the assumption that all hominid teeth from each site belonged to a particular taxon. My procedure here has been to plot the more complete dental profiles of sepa- rate individuals and to describe the types that occur. Then all possible speci- mens are compared individually and classified according to which type thei-r dental profiles most closely resemble. Obviously this cannot be complete as many specimens include just one or a few teeth and show no clear pattern. If this procedure had put all el-assifiable individuals from Swartkrans and Kromdraai in one category and all those from Sterkfontein and Makapansgat in the other, it might then have been a good guess (and no more) that all other individuals would likewise belong to the type traditionally associated with their sites. Actually the first sorting, by dental type, failed to support the accept,ed site allocati.ons. In contructing dental profiles it soon became obvious that M 3 was an erratic tooth. Its area often bore l-ittle relation to that of the preceding moIar. In many cases Lwo or more individuals with otherwise si-milar profiles differed greatly in the size of their last molars. Also there were many in- stances where the right and left ttwisdomtt teeth of the same individual were markedly different. Accordingly, this tooth was left out of any serious con- sideration ln assignments to types. The first incisors were also found to be more confusing than helpful, but in a different r,ray. In the same individuals (and those determined to be of the same type) the upper and lower dental profiles proved to be remarkably similar except for these first incisors. The tooth is markedly larger in the upper jaw. This is quite natural as its great mesiodist,al J-ength ensures that all sub- sequent upper teeth overl-ap their lower counterparts distally by half a cusp 1-ength. This results in cusp alternations which tend to keep newly erupted teeth in their proper occlusal reLationships until their roots are firmly es- tabllshed. If this first incisor i"s ignored, the terminology used to describe the shapes of the dental profiles is exactly the same for both upper and lower jaws. The various dental profiles are also compared with those of genus Homo as drawn from data conpiled by I'Iolpoff (197f).
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